SMITHSONIAN CONTRIBUTIONS TO THE EARTH SCIENCES NUMBER 6 Daniel J. Stanley Distribution of and Noel P. James EchinarachniuS p (Lamarck) and Associated Fauna on Sable Island Bank, Southeast Canada ISSUED APR 2 7 1971 SMITHSONIAN INSTITUTION PRESS CITY OF WASHINGTON 1971 ABSTRACT Stanley. Daniel J., and Noel P. James. Distribution of Echinarachnius parma (Lamarck) and Associated Fauna on Sable Island Bank, Southeast Canada. Smithsonian Contributions to the Earth Sciences, number 6, 24 pages, 1971.?A combined bottom photographic and sampling survey of Sable Island Bank south- east of Nova Scotia, Canada, reveals locally high densities (to 180 individual/m2) of the northern sand dollar Echinarachnius parma. Populations of this form are closely related to texture of the sea floor and generally concentrated on mod- erately sorted fine to medium sand surfaces. Topography and current regime are also correlatable factors; depth, time, salinity, and temperature apparently are not. Sand dollars are second in importance, after current activity, in reworking surficial sediments, and these organisms modify at least a third of the total Bank surface in the study area. Bioturbation is particularly intense in the sector north of Sable Island. Associated epifauna and infauna populations occur in two east- west trending areas on the Bank north and south of Sable Island. Absence of conspicuous fauna, save E. parma, in an east-west zone along the crest of the Bank and near Sable Island results from extremely strong current activity concentrated in this region. Official publication date is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. UNITED STATES GOVERNMENT PRINTING OFFICEWASHINGTON: 1971 For sale by the Superintendent of Documents, U.S. Government Printing OfficeWashington, D.C. 20402 - Price 45 cents Daniel J. Stanley and Noel P. James Distribution of Echinarachnius parma (Lamarck) and Associated Fauna on Sable Island Bank, Southeast Canada Introduction The present study was conducted as an outgrowth of a sedimentological survey (James and Stanley 1968) of the sand-covered, current-dominated shal- low outer banks of the Nova Scotian Shelf off southeastern Canada. It is an attempt to evaluate more precisely than heretofore possible the fauna likely to affect surficial sediments on a shallow platform, Sable Island Bank, located far from the present mainland. It is possible as a result of this investigation to define the regional distribution pattern and density of the northern sand dollar, Echinarachnius parma (Lamarck), and associated fauna commonly encountered on Sable Island Bank. Previous investigations (James 1966; James and Stanley 1968) have shown that this circumboreal echinoderm is particularly abundant on Sable Island Bank, the largest of the outer banks on the shelf. Availability of detailed geographic and hydro- logic data, together with information on the sedi- ment regimen, suggested that a study of the benthic fauna of this area would be particularly profitable. Data evaluated are derived mainly from bottom photographs, supplemented by surficial samples collected at numerous stations distributed over most of Sable Island Bank and immediately ad- Daniel J. Stanley, Division of Sedimentology, National Mu- seum of Natural History, Smithsonian Institution, Washing- ton, D.C. 20560 Noel P. James, Department of Geological Sciences, McGill University, Montreal, Quebec, Canada jacent areas. The present analyses serve, first of all, to supplement the surprisingly small amount of quantitative data and field information on this common sand dollar (Moore 1966). This echinoid occurs on both margins of northern North America, in the western North Pacific, and in the western North Atlantic the known range of E. parma ex- tends from Cape Hatteras to Labrador and Green- land (Mortensen 1948). We relate the distribution of E. parma to such parameters as time of observa- tion, depth, topography, sediment type, current regime, and associated fauna. The problem of cor- relating fauna visually observed on the bottom with that collected in relatively small bottom grabs is also considered. Geographic and Hydrologic Setting of Study Area MORPHOLOGY.?Sable Island Bank is a sand- covered platform approximately 250 km long with a maximum width of 115 km. The 90 m (50 fms) isobath defines the bank margin (Figure 1). Sable Island, at approximately 60?W longitude and 43?55'N latitude, is located on the eastern margin of the Bank, 334 km southeast of Halifax on the Nova Scotian mainland. Topography and sediments of the island, a low (relief of 1 to 20 m) east-west trending arcuate bar of sand flats and dunes about 39 km long and 1.5 km wide, have recently been described by James and Stanley (1967). Extremely gentle gradients characterize the bank top east and south of Sable Island (approximately 1 SMITHSONIAN CONTRIBUTIONS TO EARTH SCIENCES 1:880 and 1:330 respectively) as well as north (1:280) of the island. Along its steep northern edge, the Bank is bounded by the Gully Trough, a de- pression (90 to 210 m deep) characterized by hum- mocky and dissected topography. This depression curves southward around the northeast margin of the Bank and deepens to form the southeast trend- ing submarine canyon known as The Gully Canyon (Marlowe 1967). The southeast margin of the Bank is the boundary between the outer shelf and the upper continental slope. The outer margin topog- raphy and sediments have been detailed elsewhere (Stanley and Silverberg 1969). GENERAL WATER MASS PROPERTIES.?Water mass likely to affect sediments and organisms in the study area is composed of Atlantic water diluted by 20% coastal water (Hachey 1961) and covers the outer banks of the Nova Scotian Shelf, includ- ing Sable Island Bank. Between this "shelf water" and the Gulf Stream to the south lies a band of "slope water" having intermediate surface salinities and relatively low temperatures at middepth. Shelf water is generally well stratified into three layers during the summer months and less so during the winter. The upper layer may be as much as 40 fms (73 m) thick, with temperatures ranging between 5?C and 20?C and salinities less than 32%O. The intermediate layer ranges between 17 and 80 fms (31 to 146 m) in thickness, with temperatures from 0?C to 4?C and salinities between 32%o and 33.5%O. Bottom water, occupying depths of 50 to 110 fms (91 to 201 m), has a temperature range from 5?C to 8?C and salinity gradient from 33.5%O to 35%O. The intermediate layer is most important in determining the water characteristics of the outer banks including Sable Island Bank (Hachey 1961). The presence of this cold water on the shelf is due to the large volume of water transported from the east and northeast by the Labrador Current, and chilling in situ is a continuous feature, at least during part of the year. Incursions of slope water (a mixture of surface, coastal, Gulf Stream, Labra- dor Current, and upwelling deep Atlantic waters, as described by McLellan et al. 1953) onto the Nova Scotian Shelf results is salinity variations on the outer margin of Sable Island Bank ranging from 33%O to 33.5%O and temperature variations from 5?Cto20?C. CURRENTS, WAVES, AND TIDES.?The regional distri- bution of benthic organisms whose feeding and burial habits make them dependent on the texture of surficial sediments would almost certainly be controlled to a considerable degree by water-mass movement affecting Sable Island Bank. There is ample evidence that bottom currents do, in fact, modify the sedimentary cover of the bank top. A circulatory pattern centered around Sable Island displays an average rate of drift ranging from 0.11 to 0.15 knots (5.6 to 7.7 cm/sec) (Trites 1958). This surface circulation pattern varies seasonally (Bumpus and Lauzier 1965). The northeasterly and easterly drift along the outer Bank margin con- stitutes a general off-the-shelf movement and is consistent except in spring when there is a westerly drift east of Sable Island. Bottom-circulation pat- terns in the study area are modified by marked topographic features including The Gully Canyon and Sable Island. In fall and winter, when the Bank top is most likely to be affected by storms, the number of waves over 20 feet or more are reported 5% of the time (Climatological-Oceanographic A tlas for Mari- ners 1959). In spring the number and size of waves decrease (waves less than 4 feet in height account for over 45% of all waves), and in summer waves over 5 feet in height are rare. Fall and winter are times of long period, high waves from the west and northwest. In spring and summer the direction changes 90 degrees and the waves set from the open ocean to the south. Tidal currents are particularly important in determining the current circulation around Sable Island (Cameron 1965) and modifying bed forms on the shallow Bank surface. Cotidal lines, trend- ing in an ENE-WSW direction across the shelf (Sverdrup et al. 1942), suggest that the predominant tidal flow may be NW-SE. Currents do, in fact, tend to set westward (Canadian Hydrographic Service 1960). The mean range of tides on the bank is 3.7 feet. The ebb stream sets southward at 1.5 to 2 knots (77 to 103 cm/sec) on the submerged termi- nal bars of Sable Island, and the flood stream runs 0.5 knots (26 cm/sec) less northward {Canadian Hydrographic Service 1960). Terminal bars of Sable Island directly affect the flow pattern of tidal currents in the study area; the morphology of the terminal bars is, in turn, directly molded by these currents (James and Stanley 1967). NUMBER 6 SMITHSONIAN CONTRIBUTIONS TO EARTH SCIENCES TABLE 1. ?Sample stations on Sable Island Bank, Nova Scotian Shelf Sample No. Depth(meters) PositionLongitude Latitude Composition^Gravel Colour Sample Equipment Phleger core Van Veen grab Camera 1 2 1+5 67 89 1011 1213 Ik 15 1$17 18 1920 2122 23 21+<3 26 27 28 2930 31 32 27 38 39 1+0 l+l 1+2 1+3 1+1+ 1+6 1+8 50 51 52II 55 56 57 58 59 6o 61 62P 63A 360 3922 23 28 2 20 52 256 175 286 231223 63 1+ 128 132 73 1+0 187 50 52 53 20 32 1+1+ 52 72 128 120 50 38 20 16 20 26 65 139 121 170 51 29 20 15 1+1+ 90 37 90 65 37 38 22 33 38 25 36 20 18 18 59*03.0' 59*15.5' 59*23.5' 59*33-5' 59*38.2' 59*^1-5' 59*31.5' 59*2l+.O' 59*18.0' 59*07-5' 59*15.0' 59*23.0' 59*29.0' 59*37-0' 59*39.0' 59*37-5' 59*32.5' 59*26.5' 59*1+3.0' 59*1+3.0' 59*1+2.0' 59*1+2.0' 59*1+2.0' 59*1+8.0' 59*1+7.5'59*1+8.0' 59*1+7.0'59*^8.5' 59*1+8.0'sample 59*51+.0' 59*5^-0' 59*53-5' 59*5^-0' 59*5^.0'59*51+.0? 60*01.0' 6o?oi.O' 60*00.6' 6o?oi.O' 60*01.0' 60*09.0' 60*07.5' 60*07.5' 60*20.0' 60*07.0' 60*12.0' 60*17.0' 60*20.5' 60*35.0' 60*26.1+' 60*20.0' 60*17.0' 60*11.5' 60*09.5' 60*17.0' 60*32.o? 60*1+3.0* 60*1+7.0' 60*35.0' 60*27.0' 60*19.0' 60*21.0' i+i+*oo.8f i+i+*oo.5' ^3*58.5' l+i+*o6.0' i+i+*08.0' l+l+'ll+.O' l+l+*O2.5' l+i+*26.5' l+i+*28.5' . . ? ^ - ^ l+l+*O2.5' 1+3*58.0' l+l+*00.0' l+l+*02.0' l+l+*20.0' i+i+*O7.5? i+i+*O3.5' 1+3*59.0' 1+3*57.5' ^3*57.5' ^3*58.5' l+l+?10.0' l+l+*05.0' ^3*5? !o* l+l+*O2.5' l+l+?01.0' 1+3*59.0' I4V05.0' kk*09.0' 1+3*57.0' 1+3*57.2' 1+3*57.0' 1+3*57-0 1+3*51+.5' small speck mud 0.00 98.7 1.3 0.00 98.8 1.2 0.00 97.9 2.1 0.00 98.7 1.3 0.00 99.0 1.0 0.00 98.3 1.7 0.00 99.1 0.9 0.00 99.1+ 0.6 96.I+ 2.7 0.8 0.00 38.0 62.0 0.00 72.3 27.7 0.00 60.8 39.2 0,00 97.8 2.2 0.00 99/3 0.7 0.00 89.8 10.2 8.5 85.5 6.0 18.3 80.6 1.1 3-3 95-2 1.5 0.00 35.2 61+.8 0.00 91+.9 5.1 0.00 90.2 9.8 0.00 97.3 2.7 1.0 97-7 1.3 2.2 96.3 1.5 00.0 97.8 2.2 00.0 97.5 2.5 3.7 95.I 1.2 0.00 87.9 12.1 00.0 l+.l 0.7 00.0 00.0 00.0 00.0 00.0 00.0 0.710.9 00.0 00.0 1.900.0 00.0 1.7 71+.7 00.0 19.1 1.0 2.1 00.0 00.0 00.0 00,0 00.0 00.0 00.0 00.0 00.0 00.0 00.0 52.1 90.3 97.0 98.8 98.599.0 99.2 9k. k 77 5.5 2.3 1.2 1.5 1.0 0.8 1.2 1.2 1.3 3 22.0i 1 5-2k 95.6 8 1. 97 A98.5 99.097.5 21+.1 75. 97- 97. 98.7 99.098.8 99.298.9 98.798.8 99.099.0 98.9 .2 0.7 1.5 1.0 0.9 1.2 1.2 5-8 1.6 0.6 l.l 1-3 1.0 1.2 0.8 1.11.3 1.21.0 1.01.1 5Y5/2 2.5Y6.5/1+ 2.5Y6.5A2.5Y6/3 2.5Y6.5/I+ 2.5Y7/2 2-5Y7/5 2.5Y6.5/I+ 2.5Y6/I+ 5Y1+/2 5Y1+/2 5Y6/2 2.5Y 6.5/2 5Y1+/2 5Y5/1+ 5Y6/1+ 2.5Y6.5/8 5Y1+/2 5Y5-5/2 5Y5/2 5Y6/2 2.5Y6/3 5Y6/2 2.5Y6/1+ 5Y5/I+ 10YR 6/8 5Y1+/2 5Y5A5YI+/3 2.5Y6/I+2.5YU.5/I+ 5*5/3. 2.5Y6/32.5Y6/3 .5Y6/3 ?5Y6/2 .5Y6/1+ ? 5Y1+/1+ 7- 2.5Y6/1+ 2.5Y6/1+ 2.5Y6/3 2.5Y6/2 2,5Y6/i+5/ 2.5Y6/1+ 2.5Y1+/2 2.5Y5/I+ 2.5Y6/1+ 5Y5/2 2.5Y6/3 2.5Y6/1+ 2.5Y6/1+ 2.5Y6/1+ 2.5Y6/1+ 2.5Y6/1+ 2.5Y6/1+ 2.5Y6/1+ 2.5Y6/1+ 2.5Y6/1+ NUMBER 6 TABLE 1. ? Sample stations on Sable Island Bank, Nova Scotian Shelf? Continued Sample No. Depth (meters) Position Longitude Latitude Composition ^Gravel $Sand $Mud Colour Sample Equipment Phleger cure VanVeen grab Camera 6465 66 6768 69 70 71 72 73 74 7576 7778 7980 81 82 8384 8586 87 88 89 90 9192 9394 9596 9798 99 100 101 102 103 104 105 106107 108 109 110 111 112 113 114 20 10 38 40 3662 5357 36 20 10 45 59 105 175 50 40 18 18 20 27 46 92 320 54 42 37 10 22 40 46 120 440 80 60 42 40 31 235 99732 230 7654 38 26 20 24 - 102 6o'6o' 60' 60' 60' 60' 60' 60' 6oc 60* 6o?60* 60* 60* 60* 60* 60? 60' 60*60* 60* 6o?60* 59? 59? 59? 59? 59? 59? 59? 59? 59?59* 59* 59* 59? 59?59* 59? 59? 59? 59? 59?59? 59? 59? 59? 59? 59? 59? 59? '13-5' '10.0' '17.0' '31-51 '43.2' '35-O' '26.2' '18.5' '14.0' 11.5' 07.5' 12.5' 17.0* 21.0' 07-5' 08.0' 08.0' O7.O1 07-5' 01.0' 01.0' 01.0' 01.0' 53-5' 52.0' 53-5' 53-5' 55.0' 48.0' V7.51 ^7-5' 48.0' 42.0' 43.01 43.0' 42.5' 36.5; 42.5' 42.5' 29.01 30.0' 14.0' 24.5' '29-5' 34.o1 38.0' 00.0' 31.0' 16.01 '06.0' 43 57-51 43*57.0' 43*53.5' 43*50.0' 43*46.5' 43*37.0' 43*43.5' ^49,0' ]? >t ?c X, ^' 43*56.5' 43*49.5' 43*40.0' 43*30.0' 43*30.6' 43*45.0' 43*50.0' 43*54.0' 43*41.5' 43*54.5- 43*52.5' 43*49.0' 43*37.5' 43*31.5' 43*46.0' 43*49.0' 43*51.5' l|.x?SS 0' 43*52.0' 43*49.0' 43*37.3' 43*34.0' 43*41.0' 43*46.0' 43*51.0' 43*52.0' 44*55.0' 43*58.0' 43*59.5' 43*43.5' 43*37.5' ^3*40.0' 43*46.5' 43*51.5- 43*55.5' 43*57.2' 43*58.0' 43*58.0' 43*53.5' 43*50.5' 00.0 2.0 00.0 00.0 00.0 00.0 00.0 00.0 00.0 00.0 00.0 0.8 00.0 00.0 00.0 11.85.6 00.0 00.0 00.0 00.0 00.0 3.5 / 00.0 00.0 00.0 00.0 1.3 00.0 00.0 0.4 00.0 0.9 00.0 00.0 00.0 00.0 12.4 0.00 00.0 00.0 00.0 20.4 00.0 7.5 00.0 00.0 00.0 00.0 00.0 99.9 97.2 98.8 95.6 98.8 98.0 98.4 98.1 98.8 99.0 98.9 96.7 98.0 97-0 9^.5 87.3 92.8 98.8 99-0 96.4 98.5 97.3 95.3/ 97.8 96.9 98.1 98.9 97.8 98.0 97.1. 95- h 31.8 97.3 96.3 97.5 98.5 98.6 86.5 98.8 80.5 58.6 94.8 72.2 98.0 90.1 99.1 98.7 98.9 98-9 99.2 1.0 0.8 1.2 4.4 1.2 2.0 1.6 1.9 1.2 1.0 1.1 2.5 2.0 3.0 5.5 0.9 1.6 1.2 1.0 3.6 1.5 2.7 1.2 / 2.2 3.1 1.9 1.1 0.9 2.0 2.9 4.2 69.2 1.8 3-7 2.5 1.51.4 1.1 1.2 19.5 41.4 5.2 2.0 2.4 0.9 1.31.1 1.1 0.8 2.5Y7/4 10YR7/2 2.5Y6/4 2.5Y5/2 2.5Y5A 5Y5/2 5Y5/2 5Y5/2 2.5Y6/2 2.5Y6/3 5Y6/2 5Y5/2 5Y5/2 5Y5/2 5Y5/2 2.5Y6/4 5Y5/2 2.5Y7/2 2.5Y6/2 7-5Y3/2 2.5Y6/2 5Y5/2 5Y5/2 ioyR6/2 7-5Y5/4 5Y5/2 5Y5/1 5Y7/2 2.5Y6.5/4 5Y5/2 5Y5/2 2.5Y5/2 5Y5/1 2.5Y6/4 5Y6/2 5Y5/2 5Y5/2 5*5/2 2.5Y6.5A 2.5Y7/3 5*5/2 5Y5/2 1OY5/1.5 5*5-5/2 5Y6/2 5Y6/2 2.5Y7/3 2.5Y6A 2.5Y6/4 2.5Y6/6 SMITHSONIAN CONTRIBUTIONS TO EARTH SCIENCES SEDIMENT DISPERSAL.?Bottom currents mold the surface of Sable Island Bank in response to water- mass movement of the type described in the pre- vious section. Large sedimentary features, includ- ing sand waves and subaqueous dunes in water less than 40 fms (73 m) deep, show that net current movement north and south of Sable Island is in opposite directions: toward the east in the region north of the island, and toward the west, south of the island (James and Stanley 1968). Intensity of current activity increases, as would be expected, in the shallower water close to the island. This cir- culatory pattern is confirmed by an independent examination of the texture and mineralogy of the sand on the Bank. Most of the sediment on the Bank surface is now (or in the recent past) being moved by bot- tom currents. Winnowing and bypassing of finer sediment would explain the absence of fine silt and clay on the Bank surface (Stanley and Cok 1968). Although much of the Bank is undergoing active erosion, several zones of probable sediment accum- ulation are on the Bank margins (shelf-break) south, east, and northeast of Sable Island (James and Stanley 1968, their Figure 10). Methods A total of 114 bottom grab sample stations were taken on the Bank and surrounding area during the period from 5 to 9 May 1965 (Table 1 and Figure 1). The sample grid was planned so that the density of stations increases toward Sable Island. A Van Veen type grab sampler, which covers approxi- mately 0.15 m2 of surface area and when full re- covers about 20 kg of sediment, was used on 110 of these stations. Small Phleger cores were collected at 4 of the deeper stations. Camera lowerings were made at 32 of these stations with an EG&G bottom camera rig, below which was suspended at 7.6-cm- diameter compass for orientation of physical and biogenic features. The camera was automatically triggered (not bottom-activated) and photographs were taken every 12 to 15 seconds while the camera rig was about 1 to 4 meters above the sea floor. The camera rig remained near the bottom for approxi- mately 5 to 10 minutes as the ship drifted on sta- tion. The direction of drift on station is indicated by a 25-cm-long vane attached to the compass. A total of 307 (1 to 30 per station) usable black and white 35-mm photographs were obtained at 28 of the 32 camera stations. A small clock show- ing the exact time when photographs were taken and a number recording the frame sequence, noted in the data chamber, were also photographed. Depth of stations ranged from 10 to 250 meters. Position, water depth, and textural composition of bottom sediment of each sample station are listed in Table 1. Complete cruise data and results of mineralogical analyses of the bottom samples col- lected at the 114 stations are detailed in James (1966). Negatives of all photographs are main- tained in the bottom-photograph collection in the Division of Sedimentology, National Museum of Natural History. Data obtained at each of the 28 camera stations are listed in an extensive table available from the National Oceanographic Data Center, Washington, D.C. (NODC Accession num- ber 70-1010). From each usable photograph the area photographed was calculated and the follow- ing information recorded: quality of photographs, presence of compass in photo frame, presence of ripple marks, sand waves and other current indi- cators, mottling, bioturbate structures (tracks and burrows), and fauna. Where one particular species was abundant (e.g., E. parma or anemones), the total number of individuals was counted. In the case of E. parma, apparently stationary, partially buried and moving individuals were counted sepa- rately. The occasional white, often overturned, sand dollar was interpreted to be dead. Finally, the num- ber of individuals per square meter was calculated. Sediment Texture and Evidence of Current Activity Bottom photographs supplement information ob- tained from the mineralogical and textural anal- yses (Figure 5). Distinctly different types of sea- floor microtopography and texture were noted (Plate 1) : clean sand, generally rippled; shelly sand with granular texture; silt and silty sand bot- tom; and sea floor completely modified by burrow- ing organisms (bioturbation). A light and dark mottled bottom was observed at a number of sta- tions. The dark material is almost certainly or- ganic; the patchy distribution is produced by bur- rowing organisms and bottom-current activity that disrupt this bottom veneer. NUMBER 6 ? o M " *3 a . 3> cs 8 S '-* o rt rt 5 ? ? a II SJ g g & & la Illf S 5 S 8 3 i1-8 8 SMITHSONIAN CONTRIBUTIONS TO EARTH SCIENCES Bottom photographs also provide evidence of short-term bottom-current activity (Plate 2). Most useful in detailing current direction are sand rip- ples noted at 14 camera stations (Figure 2). In one instance, the predominant orientation of anem- ones, presumably bending in the current direction (Station 13), was also noted. The NW-SE trending crests at Station 82 are those of larger sand waves. The ripple-crest orientation was recorded at all 14 stations, and the sense of bottom-current move- ment determined at 8 of these (Figure 2). In shallow water surrounding Sable Island, the orientation of ripple marks confirms the circula- tory pattern described earlier. Northeast of the island, at Station 59, a NW trend off the Bank is suggested. Current directions recorded in the head of The Gully Canyon (Stations 9 and 13) indicate a movement seaward, generally parallel to the canyon axis. A predominant downslope current movement is also noted at the shelf-break near the head of Sable Island Canyon (Station 18). It is noteworthy that areas presenting evidence of strong current activity and those illustrating mottled sediment are generally mutually exclusive (Station 55 is an exception). At Station 9, what appears to be eroded algal matter has been con- centrated in ripple troughs by bottom-current ac- tivity. Echinarachnius parma on Sable Island Bank POPULATION DENSITY.?The northern sand dollar Echinarachnius parma (Lamarck) was observed in 13 of 28 camera stations. Where present, the den- sity ranged to as many as 185 individuals per square meter (at Station 21). The number of photographs showing this species varied from 5 to 22 at any one station. The photo-to-photo variation in den- sity at each station is depicted on- the histograms shown in Figure 3. Immediately apparent is the irregularity of con- centration (Plate 3) caused by clustering of in- dividuals. From comparison of the number of animals counted within the area covered by the photograph (shown in black on histograms in Figure 3), it appears that if the area covered ex- ceeds one square meter, the number counted is relatively close to the mean for that particular station (see Stations 8 and 21). Only a few photo- graphs suffice then to give a reasonably close ap- proximation to the overall distribution at any one position on the bank. It is noteworthy that correlation between individ- uals photographed at the surface and individuals retrieved in a small grab at the same station is poor (Figure 3). Those areas that contain more than 50 sand dollars per m2 yielded two or more individuals in the accompanying small Van Veen grab sample. This low retrieval reflects the small area covered by the grab but also suggests that not many sand dollars were buried at the time of the survey. REGIONAL DISTRIBUTION.?A chart showing the distribution and density of E. parma on Sable Is- land Bank is given in Figure 4. This chart is com- piled from both bottom photographs and grab samples (the presence of at least 2 individuals or more retrieved in a grab sample at a station where no photos are available indicates the probable abundance of at least 50 individuals/m2). The northern sand dollar is present over one third of the total Bank surface. Highest densities are found in an arcuate trending belt north of Sable Island as earlier postulated in a sedimentological study of the Bank (James and Stanley 1968) . Concentra- tions of up to 180 individuals per m2 are found just north of the two terminal submerged bars ex- tending from Sable Island. High densities (>20/ m2) are rarely found in water deeper than 30 fms (55 m). While a large area of relatively few sand dollars is found on the Bank southeast of Sable Island, the crests of the island's two submerged terminal bars, nearshore stations, and the area southeast of the island appear devoid of sand dollars as well. Sand dollars do occur, however, in two restricted areas southeast of the island: (a) a linear NE-SW trend in shallow water just south of Sable Island's eastern submerged bar, and (b) at depths of 100 meters on the eastern margin of the Bank adjacent to The Gully Canyon. The deepest stations where individuals are en- countered are near the heads of Sable Island Can- yon (Station 78, 175 m) and The Gully Canyon (Station 9, 256 m). A number of trends were observed in the detailed photographic analysis. Comparison of sand dollars with the known compass diameter indicates that the NUMBER 6 08:17 19:00 (5) 200 100 200 100 11:25 15:16 1 10:10 12:30 (it) 12^25 (111) ... Station number 100 Sequenc eof photo it* / E. parma retrieved in grab LEGEND FIGURE 3.?Histograms showing photo-to-photo variation in concentration of the northern sand dollar Echinarachnius parma (Lamarck) (density in m2) as observed at camera and grab stations on Sable Island Bank. The area covered in each photograph and time of sampling is also shown. 10 SMITHSONIAN CONTRIBUTIONS TO EARTH SCIENCES "a u &I ? a * .S -o O M It 18 .2 *- NUMBER 6 11 6100 30 6000 59?00 S100 5900 SABLE ISLAND BANK MEAM GRAIN SIZE (M7 ) (After Folk ond Ward , I957) z I I 3.0 VERY PINE SAND FICURE 5.?Distribution of mean grain size (in units) on Sable Island Bank (after James and Stanley 1968). Note close relation between the presence of E. parma (see Figure 4) and that of fine to medium sand grade. predominant size of E. parma observed in photos is between 5 and 6 cm. This is in close agreement with dimensions of this form found off New Eng- land (Lohavanijaya and Swan 1965). It is difficult to estimate the amount of movement of individuals in some photographs due to the absence of re- worked sand markings. In many cases, however, partially buried forms and those having left a trail are noted (Plate 4). There appears to be no relation between numbers of individuals and time of day or in the tidal cycle (Figure 3). Only in a few cases were individuals found in concentrations such as those observed by Zenkevitch (1963) in the northwest Pacific. With the possible exception of Station 56 (Plate 4) in- dividuals appear to be moving in a random pattern, confirming the observations of Weihe and Gray (1968) and Boehmer (1970) on southern United States species of sand dollars. In areas of high den- sity this movement is responsible for modifying physically produced bottom structures, and the effect of this bioturbation can be seen on Plate 4. The movement, by rotation and progression, as described by Parker (1927) in response to feeding and burying, modifies both rippled and nonrippled surfaces. The broad, flat, generally arcuate tracks made by the sand dollars are characteristic and may be recognized in the fossil record. At Station 9 (256 m) located in the head of The Gully Canyon, sand dollars were observed on a rippled sand bottom in which organic matter, prob- ably algal detritus, was noted in ripple troughs (Plate 2f). Other Epifauna In addition to E. parma the presence of other or- ganisms living on the sea floor as noted in bottom camera stations and grab recoveries include star- fish, brittle stars, anemones, gastropods, and fish (hake and cod). Their distribution is shown in Figure 6 and examples are illustrated in Plate 5. Starfish [Henricia sanguinolenta (O. F. Miiller) and others] are found at 8 camera stations from depths of 20 to over 200 m and appear more nu- merous north of Sable Island. Brittle stars (Ophi- 12 SMITHSONIAN CONTRIBUTIONS TO EARTH SCIENCES ? 5 ? S o c a, >~ .2 a, NUMBER 6 13 -s w O ^ s Hl e3 o 3| ?1 > 2 am , "3 18 So .2 3 o 14 6100 6000 SMITHSONIAN CONTRIBUTIONS TO EARTH SCIENCES 30 5S[00 61 00 5900 SABLE ISLAND BANK SORTING (After Folk and Word,1957) I I <0.35 V.WELL 0.35-0.5 WELL SORTED 0.5 - 0.71 MODERATar WELL >0.71 MODERATELY FIGURE 8.?Regional distribution of sorting of sand on Sable Island Bank. The zones generally devoid of fauna north and south of Sable Island are very well sorted sands coinciding with areas of high current activity (Figure 2). omusium lymani Wyville Thomson at 2 camera stations and in 3 grab samples), on the other hand, were not recorded above 90 m. These ophiuroids appear not to be restricted geographically but are rather limited to deeper water (for example, Gully Trough, The Gully Canyon, and outer shelf near the shelfbreak). Numerous attached anemones were observed at 6 deep water camera stations (>90 m); north of Sable Island, at Station 33 (50 m), several ane- mones were attached to shells. At Station 51 in the Gully Trough (depth of 146 m) as many as 98 (26/m2) were counted in a single photograph. Gastropods, including the species Aporrhais oc- cidentalis, were recognized at 8 camera stations and recovered in 7 grab samples. These mollusks are more common north of Sable Island. Gastropods occur in water deeper than 55 m on the southern part of the Bank, while north of Sable Island they are found in depths over 35 m and in even shal- lower water in the lee of the submerged terminal bars of the island. The organisms cited above generally occur to- gether. The regional distribution of this epifaunal assemblage occurs in east-west trending zones as depicted in Figure 4. A belt 34 to 40 km wide that parallels the shallow crestal axis of the Bank is generally devoid of conspicuous epifauna. Infauna The regional distribution of organisms living within the surficial sediments cover of Sable Is- land Bank, evaluated from both photographs and grab recoveries, is shown in Figure 7. Included in this assemblage are tubes and polychaetes from grab recoveries and burrows and tubes observed in photos. Actual examples are shown in Plate 6. The regional distribution pattern of this assem- blage is similar to that of the epifauna (Figure 6). The zone devoid of infauna, however, is nar- rower (12 to 22 km) than that devoid of epifauna and is more symmetrically oriented along the crest of the Bank. Infauna is generally associated with a mottled sediment surface. Small mounds of light-colored NUMBER 6 15 sand surrounding burrow openings (Plate 6) are separated by darker areas, suggesting that this patchy coloring results, in some cases, from bio- turbation (i.e., disruption of an originally con- tinuous dark veneer). Recognizable pelecypod shells, including Cyrto- daria siliqua, Pecten magellanicus, and Arctica islandica, are observed in bottom photographs. Fragments of shell, or shell hash, are ubiquitous. Bottom photographs are generally unreliable in estimating the density of pelecypods because of the large number of buried individuals. Bottom grab samples reveal a concentration of shell fragments in southeast, northeast, and northwest belts trend- ing away from Sable Island toward deeper water on the Bank (in James and Stanley 1968, their Figure 6B) . Shells are also abundant near the south shore of the island, at ends of both bars, and in shallow water surrounding submerged offshore terminal bars. Discussion INTERPRETATION OF E. parma DISTRIBUTION.? There does not appear to be any recognizable cor- relation between density and regional distribution of E. parma and time of observation or depth of water on the Bank. The tolerance of this species to significantly large seasonal variations of temp- erature and salinities cited earlier is apparent. The salinity and temperature of bottom water north and south of Sable Island are not significantly different and cannot explain the remarkable faunal differ- ence in these two regions (i.e., extremely large number of sand dollars north of the island as op- posed to low densities south of the island). Preda- tors that may affect regional distribution are un- known. The distribution pattern of the northern sand dollar (Figure 4) can, on the other hand, be closely related to that of the mean grain size of sand (Figure 5) which in turn reflects local condi- tions of bottom current agitation and topography. The most abundant faunas appear in the fine (2.0 to 3.0 ) and medium (1.0 to 2.0